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HIERARCHIAL BEHAVIOR IN THE 
SOUTH AFRICAN CLAWED FROG, 
Xenopus laevis Daudin 



ROBERT RICE HAUBRICH 



A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF 

THE UNIVERSITY OF FLORIDA 
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE 
DEGREE OF DOCTOR OF PHILOSOPHY 



UNIVERSITY OF FLORIDA 
August, 1957 



She vrl-ter vlshes especially to aekncnrled^ Ms ladebtedaess to the 
late Dr. W. C. Allee and his vlfe, Mrs. W. C. Allee (Ann Silver), uhose ideas, 
encourageiaent aad advice vere Instrumental in the productioEi of this vork. 
fhe close personal relationship with both of than remains one of his most 
cherished oeBories. 

She writer also vlshes to Idiank Sr. B. Soffin Jbnes for taking on 
the hurdsn of this program after the death of Dr. Allee, and vlshes to 
adsxovled^ his indebtedness to him for eneouragem«nt and suggestions. 

Qratitude also goes out to Dr. H. M. Uallbrunn -vho gave advice on 
statistics. Dr. C. J. Gk>in vho suggested the e3q;>erltDental animal, Drs. L* 

* 

Bemer, J. H. Gregg, R. H. Waters and S. S. Wiaiberly vho criticized the 
text and to many fellow studaats Including C. D. Wilder, W. C. Sloan, C. D. 
Hynes, J. f . Howill and others for their discussions, critlclams and ideas 
relating to the developnent of this vork. 



11 



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BIOCBRAPHKAL I3B8 



INTROIUCTIOR 



Hlerarchlal b«faavlor is a phenoaeam In indlvldu&ls display 

certain aggressive actlcns toward one another such as pecking or nipping. 
Differences in the frequency of these actions are often consistent enou^ 
to enable cue to rank individuals as to aggressive potential or position 
in relation to each other. Bvidaace of hiexwrcMal behavior Is fomd la 
such divergent parts of the animal kingdom as Arthropoda and Oiordata. 

Among invertebrates such behavior has been reported by Allee and 
Douglls (191*5) in the hermit crab, Pagurus longLcarpus. by Dou^s (l^) 
In the lobster, Hoaarus anerlcanus, by Bardli {19^) in the Tiasp, Pollstes 
galllGus, by Bwrjberg (1953) in the crayfish, Orooectes vinUs and by 
Lowe (1956) in the cre^yflah, Caaibarell\is shufeldtii. 

Aaong vertebrates evidence of hieararchial behavior is abundant in 
the class Osteichthyes. Braddock (19^5) demanstrated it In Platypqeellna 
aaculatus, as did Allee, Greenberg, Rosenthal and fraxik {19^) in the gireai 
eunfish, Leponis cyanelltxs . Hewnan's (195^) work gives infonnation on intrar 
and interspecific conrpetltion as well as hlerarchlal behavior in two trout 
■pedes, aaloelinns fontinalls (Mitchell) and Salao gairdaeri Richardson. 
Evidence of hlerarchlal behavior in the class Reptilla was reported by 
CSreenberg and Noble (19^*) in the American CSxameleon, Anolls carolinensis. 

Members of the class Aves probably have been studied with the 
greatest frequency and have yielded most in the vay of facts and principles 
of hlerarchlal behavior. Papers by Schjeldenq>-a!bbe (1913* 1935) aad Quhl 



and AllM (X^) on th« dooestic tcnrl, Eeuoks (195^) m ,}tBigIe fowl, 
Mtibs^hlag (19^1) on plgeoas, B«mett (1939) on ring dores, Hovard and 
Anlen (19^2) on <|uall, HauBaerstram (l9'^2) on chickadees and Sabine (19^) 
m JtDieoa and tree aparrova Indicate the vide cov«M0t in -^a elasa on 
Tjoth dame sti Gated and vild forms. . .j, . . 

Wbrk on rats "by Uhrich (1938)^ aqolrrels "by Gordon (l9kO), infra- 
htanaa primates by Maslov (I9U0), chlarpanssees by Crawford (19^2) and Yerkea 
(19^ ^ Rhesus laanlaay by Carpenter (19^2) all demonstrate the social 
hlerardiy in the class MaBiaalla* A form of hlerarehial organlssatlaa In 
htman Indlviaaal and grottp l)rtucvlar Is Indicated by HasMns (1951)* Allee 
(1951), Keith (l9*^9), Sherlf (1956) and other*. 

Hhile 'Uiis phenotaenon reaches Its grtfttwit eotoplexlty aoong verte- 
brates (Allee, 1952) mtll recaitly no hlerarcMal behacslor had be«i re- 
corded far the classes Agoatha, Chondrlchthyes or An^pliibla. Bie first breajt 
111 this Bttuation occiirred vhen Allee and lUcldLnaon (l95*^) reported finding 
a primitive form of this behavior in the dogfish ehark, Masteliis eaalSi a 
member of the class Chmdrlehthyes. !Qais leaves A^gaa-ttxa, regarded as the 
most primitive vertebrate daaa, «aA ih^phlbla, a claaa of medlm coBq?lexlty 
as the oaaly two elaaaes of Vertebrates in which there has been no report of 
hlerarehial b^unrior, 

Prtfvious works have Indicated aggresslcn end territoriality in tSie 
class Aatphibia, but not hlerarehial behavior. Martof (1953) reported on a 
sort of territoriality observed among breeding males of the gre«i tros, Bana 
clamltapis, based on Isolation, a type of intolerance, advertiseaoent and 
fixation. He augcgeated tbat a relatively aggreaaive perioa is followed by 



one of stability, lAloh ColllaB (19^) cmsldered to b« the case In oth«r 
vertebrates. Pearson (19^^) found in the spadefoot toad, Scaphiopus 
boIbroQki holbroolgl (Harlan), avoidance reactions smoag Individuals and a 
type of tmdef ended home site territory vhlch seldom <9rerlapped others even 
in crowded areas. ^Test (195^) reported the first definite case of aggresslca 
In the class Auzphlhla as displayed iA dtffnat o£ ttzxitarjr "by the frog, 
Phylobates trinltatis Boulaiger. Bvldence of territoriality In the bull* 
frog, Rana catesbelaoa, was reported by 0. B. Ctoln (1955 )• ^ ^& vas 
observed to croak and jm^ at a smaller frog In the same general area vhlch 
resulted in the retreat of the smaller animal. Jn addition, other indi- 
viduals vere consistently fo^{nd in certain areas located around a pool* 
Qrant (l955) also reported on a case of territoriality in two species of 
salamanders, Eurycea bislineatja and Hemidactyllum scutatua, based upon a 
#»haniytep end a defended territory, tlie smm of defense i ncl u ded direct 
biting attack in B. bisllneata and challenge only in H. scutatum. 

Preliminary experlofints were attempted on certain species of frog« 
but the problaa of classifying the actions of the animals yblle under obser- 
vation was difficult. Use of Xenopus laevis Daiadin as the e^qperimentzQ. 
animal was suggested by C. J. Goin (1953) of Ifalversity of Florida 
because of its habits and mode of life. !Ilhe purpose of this study was four- 
fold: to determine ^diether hierarchial behavior was present in Xenopus, to 
analyze the phencnKUXi of kLeocarchlal behsvloor, to fill a gap in r«qi>oart8 of 
hierarchial b^vlor In dlfflerent ph^ogeneHo groops and to add to 13m» 
information on aggression with possible applications to human behavior. 



MfcterlaXs and Methods 

Frogs tised in ^bi» study weare obtained from the sttpoply house of Jay 
E. Cook, CoclmeyBvllle, Maaryland and were received as adults by Cook from 
South Af^ca (see Walker, 19^2)* Parlor to the observatioa period animla 
vere kept in five gallon aqpiaria filled with water to a depth of ten inches. 
Ihey vere kept out of direct sunlight and t«iiperatures vere maintained 
between 6cPand 75° F. Bie vater was changed after each feeding or vh«i it 
became doudy. At first city water which had been conditioned by four days 
storage was xised, but after sane miwAl » died Xooal spring <xc stream water 
was used. Bie above methods for maintaining adults are discussed in detail 
by Parker, Bobbins and Loveridge (19^7)« - * " • 

3Eenoms laevis was first described by Daudin (1803) and is primarily 
an aqpmtic frog. Adaptations to this a^tic habit are evident in the large 
webbed >'^r'H feet that peimlt rapid B»veineat in any direction, a well-developed 
lateral line systeni (Slkan and Ifurray, 19^ araionia excretion in adult 
(Muaro, 1953), and nasal cavities lined with sensory epithelixm that functions 
In olfsotozy perception ^en submerged (Bat«r«oa^ 19$l)* ^ spite of the 
aquatic habit at times the is seen ntcnrlng orerlmd or is found In clods 

of BWd near dried up ponds (Leslie, I89O; Rose, 1950). 

Two groups of twelve female frogs each were used in this study and 
were kept in 8 - 10 inches of water at all times. ObservBtions on frogs 
1 > 12, desigoated as Ebgoeriment I or Ckroup I took place in the Marine Bio- 
logical Laboratory at Wbods Hole from June to August 195^* Observations on 



frogB ion - 112, desismted as Sxperiment II or Oroizp II, were coorpleted duiing 
the smmr aad fall of 1955 ^'t the University of Floricla. - . 

. Ibqperlinent I 

ftLx frogs iwre kei^t betwMSi observatioaa la MOh of two tea-galloa 
aquaria and vere wlsShed once a veek duriag the eaperiaeatal period (Table l). 
When animals were fir»t received certain digits on the froait limbs were clipped 
to facilitate reeogaltiott. Further clipping vas unnecessary, even thou^ 
digits grev l>ack, ■beca\i8e characteristics like size, color pattern, behavior 
and body shape varied nldaly anong iadlvidualB and awds it easy to recognise 
then alter they had been observed for a time. 

Pairs vere observed separately in five -gallon aquaria. Before each 
series of pair observations vater -was changed and t«fo different animal s vere 
used. During all observations cxibes of liver, about l/h Inch cn a side, vere 
plB«ed in aquaria oae at a tine as the animals ate them. Feeding occurred 
cBily during time of d b s e r r ation. 

^ . Activity differ«ices aiaong individuals were noted by pairing frogs In 
all possible combinaticnB. Each pair vas oibaarved a total of 200 ainutes 
divided Into ten twenty minute periods. Biese ofbserration periods nwre con- 
ducted in two sets of five each. Periods vithin each set followed each other 
consecutively, excepting short breaks between periods for lib* «aiperlmenter to 
rest, but the two sets Involving one pair of animals vere not run consecutively. 
3Dbie followlag infoimation vas recorded during the pair observations for each 

1. Hips given. A nip vas recorded when one frog nipped or bit anotiier 
and caused the other to retreat* 



2* Pushes glvei. A puBh was recorded vhea oaae frog appcroacbed another 
and displaced It throu^ contact. ^ 

3. J^^oaches* Ifunber of times each frog approached other aotloaLess 
rwBiber of pair aad touched -without further activity. 

k, I&toowa. Number of times the two aalaals made contact, vhlle in 
maticn, vlth uo dbserTable dlffereatial betwem th«BU 

5. Cuib«B of liver eaten "by eaeih ftfog and th* ordBr in iiblch each was 
obtained. 

6. !Ehe troQ obtaining food first at the start Of tioh 8«ri«8. 

Activity differences were also noted vhaa each frog was isolated a 
total of 200 minutes ^ttvlded as in liie paired sltuatlcn into two series of 
ten twenty-Bdnute periods each. Olhe following infonaatiOBi was recorded for 
each firog while isolatedi 

1. KtBober of cttbes of liver satsn^ : 

2. Hoe in seconds talaen to obtain each piece of liver. 

. ■ Ejqperlment IZ ■ 

2h IMs eaqperlment, in order to provide better control and prevent 
possible carryover of hierarchial behavior, the frogs were kept isolated in 
OM ffallGBi jttrt bstuMR obMOvations. i^alred and isolated animals were ob« 
sttred using nsthods similar to tluMM in Ixperlment Z. Temperatures varied 
between 71° - 7^° ?. and the aniioals were welgjxed weekly during pair obser- 
vatioas (OUile lA). In Experiosnt ZZ^ to dAterain* ^-Oisr poeitlcn in the 
hierarchy was consist^t under different eondltlQaB, observatlaDis wer* asds 
on sach "Tnima^ vhen in a groig> of twelve as well as \dien in a pair cooiblnatiai. 



Since it vae li^posslble to record the activities of all the frogs in the 
aguariuffi simultaneously the behavior of eaeth frog with reference to the 
other eleven vas observed for a total of 200 minutes. Each flrog "was mdnr 
ohservatlcEi for a total og 200 wln^iteii 9oA OfHif •ffbf.vities involving the 
tingle frog "being observed at -ttie moBtent irere reecaraed. For instance, no 
record was kept of idiieh animals were nipped by or received nips from the 
frog under observation. Observations vere made like those on the paired 
situation, namely, in ten i20HBinute periods at the rate of five periods a 
(laar* 3he animals vere laolated a mtnlTiniin of twelve hours after each dally 
observation • 

•- •• • ■• ..■ '. • ■■ ' ' ■ ■• - 

Besaltg 

' ' febles 1 and lA indicate restdts of weekly weijjiings in number of 
grams. Individuals in Group I varied in average veldts frca ^0 to 168 
gnats, while individuals in Group II varied frcoi 37 to lUo grazas. All of 
Qratx^ I (frogs 1 • 12) showed some gEdn during the experim^ital period as 
did all but three frogs in Gtck^ II (frogs 101 - 112). 

Vhen animals were placed together in pairs one of four different 
tjGPes of ac-U.vity wu <^s«rved9 1. e. the nip, puah, approach or unknown. 
These types of acttrlty ftccrlng emtacts wee recorded for each pairing and 
the results are shown in t&ables 2 throii^ 5« 

Besults in the nip category are recorded in Sable 2 for frogs 1 - 12, 
and in liable 2A for frogs 101 • 112. Animals were ranked in order of total 
nips given in all pairings. In Baperlment I this varied from UQ to 279; 
while In Experim^t II it varied from 3 to ^77* lianklng of frogs for total 
nips givei (totals along the bottom row) was correlated to ranking for total 



nips recelred (totals along the left coUna}* Frogs X • XS v»ri«d in total 
maaber of nips received from 76 to S^h. Hfhm. individual reoikl&gs in naaber 
of nips given were cooipared to ranldngs in nuiaber of nips received the 
results showed a ccarelation of -O.96. Frogs 101 - 112 varied in total 
maiber of nips received dftom 57 to iSk, and -when individual rankings in 
nuniber of nips glwi veare compered to twnliingB In nips received the result 
was a correlation -<J.50. A statistically significant differeice at the 
5 per c&xt level in nuniber of nips giv«i for any particular pairing is 
Indicated in the tables "by an asterisk following the greater value. 3a 
T&hle 2, 28 of the 66 pairings showed such a difference while In Kble 2A, 
3^ of the 66 showed this differaoLce. 

ItfbleB 3 and 3A axe similar in foim to the prewlous tahles "but daal 
with the ^ish category. Individuals in Eaperioent I varied in total nuniber 
Of poshes given fxm 85 to 5H. idxile those in Experla«t II varied in this 
category from 20 to Wf» Also, IndlvlAials in Qroop 1 varied in total 
pushes received trm. IO3 to 381, ^ftiile frogs in the other group varied frca 
71 to 262, When individual rankings in nunbtar 9f yoabas given were contpared 
to rankings in number of pushes received in RjqperliBent I Uie result was a 
eoarrelation of -O.87. In 'Eaqperijma.t II when rankings were cooipared the 
result was a correlation of -O.76. In Table 3, 38 of the 66 pairings showed 
statistically sigaificant differences between individuals in number of pushes 
glT«x (shown by an astarlBk fcOlowing the Xarsar total) while In Sable 3A, 
30 Of the 66 palHngii showed mudh «ffta?«nee8, . 

fobles k and kk deal with the approach category and are similar to 
those described above. Individuals in Esi^rimaBt X varied in total woiber of 



approaches made frm. klO to l^S vhile those in Bs^terlmeat II varied In this 
category from l80 to 767 ♦ Ihdlvlchials in Group I varied in the nuaiber of 
times each vas airproached £rcm a minimum of 1^23 to a wndniinn of 1207; -while 
animals in Oroup n v«rle4 from 277 "to 773 in same category. When individ- 
ual ranMngs In nuniber of approaches made vere conipea:>ed to ra nk i n gs in appro- 
aches received in Experiment X tdiae result vas a ccnrrelatioaa of -0*83, and in 
Ibqperimesat II the same ccmpartsons produced a correlation of -0.88. Id. 
Tfehle k, 1*2 of the 66 pairings showed statistically sigiificant differences, 
indicated "by asterisks, "betWMn individuals in nianber of approaches made, 
while in Table kA, 32 of the 66 pairings showed this* l . - ... 

SHe untoaoua category, represented hy Tables ^ and ^ indicated the 
otBlber of ttms trofft nade cmtacts that -were of equal or of no aggressive 
potmtial as nearly as the observer could determine. The nttmbers of con- 
taots vere the same for each meciber of each pair heoauae both Bniml . s vere 
given credit vhaiever a ccaatact of this type occurred. Tbe total nvoiber of 
ccmtacts in the unknown category among individuals varied from 667 to 2525 
in Sscperlment I and from 511 to 2li-ll in Experiment II« 

■..^> In Tables 9 and 9A total food consUKcption, as measured by cubes of 
liver eatea, is recorded far each frog in each pair combination. Ihe animals 
vere rsnked according to total ntaaber of evOftes eaten ov«r all pairings. 
Sable 9 gives results for frogs 1-12 and shows an individual variation in 
food consuaQytian from I5I to 327 unite. ISible 9Jl gtves results for frogs 
101 - 112 and the food ccnstmiptlan in tMs group varied fjram 75 to 803 units. 
When the rankings in total amount of food consuoed by the other eleven fjrogs 
labile paired vlth each frog vere compared to rankings in total amount of 



too^ eaten "by each £rog^ correlatloas q£ -0*62 atid •<Q*^ Wir* recorded, 
respectively, in Sxperimaits I aad 11. A statistically «lgn±ficaiit diff- 
erence "betveea £rogs ija azaouat of food ^ten in any particxilar pairing vas 
indicated in TIables 9 and 9A "by an asterisk follovlng the greater valiM. 
In !£able 9, 19 out of the 6C pairings showed such a difference. In Table 
9A, 26 out of the 66 pairings shovtd this difference* ''^ ' 

Table 6 indbLoates nuadber of alps azid ptu^es that vbm given and re- 
ceived by each anlTn&l vhile in a group of twelve. A total of 313 nips were 
given by the 12 frogs during -tiie observation period and Individuals varied 
in total number of nips given from 0 to I68. A total of 193 nips vere re- 
ceived by the 12 frogs during the period of observation, and individuals 
varied in the total nxaatoer of nips received flrom 7 to 36. 2he total number 
of nips given by the group did not necessarily eqpial the total received 
beoauae each f^ vas oS^aerved singly for 200 ni1nut<ea and only nips givan 
oar rtteeived by that mSaal wv reeorAed during this pariod. Bo reeorA iMui 
kept of ^dilch ftogs nipped or were nipped "bj the frog under observation. A 
negative correlatioa of O.k^ was found between total number of nips given 
amaag the 12 Individuals and total nips received by these same individuals. 
' ' .. A total of 537 pushes were given by the group of 12 frogs and 
individuals varied froci 5 to I36 in this categcury. A total of 735 pushes 
vere received by all troga and individuals varied f^rcci 19 to 1^2. Sie total 
number of pushes given did not equal the total ntanber of pushes reoaived for 
the reasooB given above. A ne:3Eitive correlation of 0.62 was found t u tt w e n 
total number of pushes glvaa among the 12 individuals and total pushes ra- 
calved by these aame individuals. !Zhe total ntmiber of nips (given and 



ree«iv«A) vas 506 uhlle the total niaober of pushes glreii emd received vas 

1272. • ■ ■ " - ■ ■• 

Saibls 7 soBMrlEes totsLU for tbe differeat lac^vlaraJL c«t«0orle8 
ftfoa the paired sltuaticai. Freqaaicies of the categories Increased em- 
sisteatly in both ea^perliaKits from a minimum at the nip to a maxtnnin at th« 
unknown. Itable 8, nblch suBoarizes ftrequencies of nips and pushes for groups 
of 12, also indicates an Increase from nip to push (506 nips, 1272 pushes). 
SUal«s 7 and 3 indicate that an inverse relationship e^sts 'betveen £re^[uency 
and aggressive iateaslty 0^ oontacts* 

CoBiparisms in total amount of food consumed by frogs -when isolated, 
paired and in groups of 12 are shoun in Seblea 10 and lOJL, She total amount 
of food eaten by all frogs in 200 minutes Increased frm Mil ttoits \diile 
isolated to UQk ^dien paired to 1199 vh'^ groups of 12. Ihe latter figure 
is glgnlficantly lar^ than iiie other two valxws (P *<0«01 end<O.Of>, 
respectively). ■ 

The number of times each f^g obtained food first at the start of a 
series of observations wts rtcorded in OSable 11. la the paired situation 
individuals varied from 4 to I6 in this category. When the group of twelve 
animals vera compered a variation £raa 0 to U nas observed. 

Individual variation in av«rage anount of tine taken to eOytalA food 
•while isolated was recorded in teble 12. !I5ie frogs in Bxperiment I varied 
txaa, 72 to 768 seconds vhile those animals in Esperlment II which ate varied 
from to 13^ seconds, but two of the animals in Experiment H ate no food 
idiiXe Isolated. 

'QBable 13 shows rankings in each of the categories for frogs 1-12 



These rankiass to be lised in correlatioai tests (Table l^i-) were obtained frcm 



the following tables! 

Hips vhile pair«d • . . . • Tables 2, 2k 

Poshes ^^lils paired * fables 3j 3^ 

Appcroaches -while paired Tables k, kk 

UaJfflown wMle paired ..*••••«»«* Sables 5> 3k 

Etps while in groups of twelve ««•••* Table 6 ..... 

Pushes -while in fiewtps of -tarelve ..... Table 6 - 

Awage -wei^t •»•••••••••••• Tables 1, lA 

Food consump-tion -lAile in pairs Tables 9t 9k 

Urn taken -to locate f ood vhile Isola-bed . Table 12 



Table ik shovs the c<nTelR-tioa aaong all of the categories ranlaed in 
T&blfi 13 for both se-ts of frogs. A measure of the sta-tistical siffiificence 
of -these values also is indicated. - . 



^^^^^^^^^^^ 

MBELOSBf IN GRAMS, MEASQRBD WKSSG EXFERIMERZAL EERIOD 
FB0G3 1-12, EXFERIMSHT I (19^1^) 



Wel^t. In Grams « Trog Ruaiber 



Bate 


1 


2 


3 




5 


6 


7 


8 


9 


10 


u 


12 


30 Jua 


131 


170 


105 


• » • 


90 


66 


• • 


66 


« • 


30 




k2 


6 Jul 


128 


168 


117 




96 


69 


* « 


70 


• * 


k6 


kl 


kk 


13 Jul 


127 


172 


109 


* • • 


98 


70 


• • 


67 


• • 


k6 


hi 


kl 


20 Jul 


127 


259 


l0l^ 


128 


95 


68 


72 


72 


53 


k6 


50 


50 


27 Jul 


135 


168 


112 


131* 


93 


73 


77 


71 


5^ 


k& 






3 Aug 


136 


172 


107 


137 


97 


69 


77 


67 




50 


55 


51 


10 Aug 


ikz 


168 


109 


iin 


101 


75 


82 


69 


58 


56 


61 


57 


17 Aug 


Ikl 


167 


113 




lOl^ 


78 


85 


72 


a 


58 


69 


a 


2k Aug 


135 


168 


112 




99 


78 


86 


68 




56 


67 


57 


30 Aug 




Vjk 


112 


137 


103 


76 


Bk 


70 


67 


63 


70 


60 


kr, Vt, 


13h 


168 


110 


138 


98 


72 


80 


69 


59 


50 


56 


52 



XABI£ U 

HEZCET, IS G»AMS, MEASURED SUBIKG PAIR OBSERVAZZOSS 
FROGS 101 - 112, EXPERIMENT U (1955) 





101 


102 


103 


10k 


105 


106 


107 


108 


109 


no 


in 


112 


19 Jun 


105 


96 


132 


ek 


7k 


6k 


65 




k2 


38 


• • 




SSS Jun 


103 


101 


136 


82 


82 


66 


68 




k3 


kl 


• * 


i 


3 Jul 


101 


106 


133 


83 


77 


60 


68 




k3 


ko 


• • 




10 Jul 


lOl* 


no 


1^3 


82 


75 


66 


78 




k3 


39 


• • 




17 Jul 


101 


108 


139 


93 


77 


68 


78 




k2 


36 


• • 


2k Jul 


98 


106 


138 


93 


78 


62 


76 




kk 


g 


• • 


k9 


31 Jul 


102 


no 


138 


90 


75 


62 


62 


79 


k9 


36 


55 


51 


7 Aug 


98 


116 


138 


lOl^ 


79 


62 


79 


81 


kl 


33 


• • 


ka 


Ik Aug 


99 


120 


ll»6 


96 


79 


71 


85 


85 


k5 


33 






7 Sep 


101 


128 


160 


86 


83 


65 


82 


85 


k2 


31 


58 




JEv* NX* 


101 


no 


ll»0 


89 


78 




76 


82 


kl 


37 


56 


kk 



UN H o\H otflO «^ir\ 



Ql tr>CO i»t r4 b- 1- 0\ t» * 01 tn 

* u\ 



coco t-vo rooivo-* •<*?o. ^ 



• H H 



irwO trxirv t— OS 



VO J* ONJt O • ONCO 
H • 



*. t 



• H H H W H 



57 



H-sf 01 >o iiNO| t<-^ o\oo (n 



5 . 

I 



15 



1| 

a 

I 



5 



I 
1 



n 



I 



g " 



I 

. f ft 



<3 X O ^ fl^ 



t 



I « t 



t- ^-^J> t>-^ »^ vo CU trv O O 
u>v£) H <j\cb ON <5 t«-vq if\ on 

H r4 H fi r4 H ri 



OHOOOOO^HOO • UN 



OOHH«nOOOOO • r* VO 



H H H 01 01 to<n^ H ^01 O 



OIMC&NOVO 'VO 



* H 



t-ON ijJgWOJgOjgJ^O 



1^ f4 p*4 <^ ^"1 •'"4 1^ 



5 . •; 

S| ' 

-I 

is 

8 1 : 

ft 
I si 



I 



■ 



I 
If 



I 



I I I 

JO 0 - 



I 



0\ 



?1 



H i-l H i-l 1-4 • CO 



^4 H H CM * 



vo 01 «>• ON r|j ij^***;:} 

#*4 



iA<no\0\Oco 



H 



CVJ 



H ^ Oi r-j irvVO 0| fO t-OO ON o. 



o _ 



ta 



i o 



I 



II 8 

** ID 



o 



a 



u 



I 



I I I 
.o 041 



17 



01 coco \o t~ I— W 

POCO CO O r-t O ^ CO ^ 

HHHCUHCUHHW 



OOHOOmHO OJ^ ON • 



: 8 



3 
3 



I 



2 



H H H r-t CU H UN .^<»J ^ 



HOi'HHCVJH H ^ 



CU H VO •00 tl\ J* _ . 
H H H • «OH H ^ CVi 



8 



I 



hhhhhhhhShhh 



1^ 



1^ 



^5 



■i 



4i 

p o« 

|g| « 
IH I 

Ml 

SI 



9^ 



o 



5§ 



51 



I I i 



a 



3 



I 



O C^ O On 
^ if\Jsp r- 



* H 



5l :iftl^^l(8^ll;^S; 15 



•« UNO 

. 3- !>-^ ^ t- ITv CO v5 



ra CO 



I 



« a >• 
o o a 

5 



^1 



« § 



•d 

IP 

S ^ 



2l ^ 



o 



1 



I 



I I I 



1 



I 



2 



HHHWH H * H 



^ *v 5i '^^ 'lOt-itnH tn 



UMfWO 



o 



TABES 6 



mSSBBR OF NIPS AUS posses giver ASD KECEZVED BI each mXi VSES 



Frog 



Nips 
QivmL Pteoeived 



Pushes 
CbLven Secttlv*cl 



Rtps-Pushes 
Giveu 



nps-Pushtts 

Becelved 



101 
102 
103 
10^ 
105 
106 
107 
108 
109 

no 
111 

112 
Sbtal 



13 


39 




12 




a6 


7 




19 




35 


8 




53 


19 






13 




38 


Ik 




89 


33 




1^ 


36 






Ik 






5 




19 


193» 


' ' 537* 


735* 



58 
19»t 
30«* 

g 

ll 

850 



''.5 



iff 



928 



# • Total nips or pushes given in this case do not necessarily ecfoal total 
I nips or pushes received laecause in the grouped sitoEition fro^s were 
observed one at a time. Ho record vas kept of vhicb frogs gave nips 
or pushes to or received aipe or puahea ixoa. tJie particular aninal 
being observed. 

Ck»rx«latlax (r^y) b«tw»ca nuogibcr o^* aijpfl given and nips received vas -0,k3 
Onnrtlatlon (r^y) between mafter of poaibes etvea txdt poshes received vas -0.62 



SABXS 1 

TREQMEHCX OF EACH CQRZACT TXPE WBEIT fROGS WERK IS PAIRS 



Hugiber Observed Per Cent of Total 
Type of Cmtact Experiment E:!iperlment 

I n I EC 



Mp 
Push 

Approach 


1552 
2691 

9033 
lllM 


im 
1890 
5239 
9527 


6A 

37.0 

>5.6 


28.9 
52.5 


mai 


^ik20 


laiuo 


100.0 


100.0 



■ V. 




naftlMJi OF EACH C(MEACT TZfE WHEN FROGS WERE IN GRCXJP OF THELVS 



lype of Ckmtact JSmSoer Observe 



.■ Hip 


506 


Ptuh 


1272 


Ibtal 


1778 



CNON 



HH H H H H H CW • « <n 0\ 

^1 



9 



i| 
11 

|l 

I'd II 



I III 

4 V* 



«n cnco £^ irv o ift^ inj* • i*\ 



co *n cu o\eo »»i t- 0\ if\ H ♦ ro co 
H HHH H*H r-j 



• to 



• 00 



I 



11 

«1 



|1 



I 

a5 



I III 



•EABIE 10 

TOTAL rarras op tood EAaaar by frogs dukekj oBSERVATioaf time of ^ 

200 HINUTiSS VSSS IS0IAS3ED, PAIRED* AND ZH CSOUP OF TMELVE 



Frog 



Total l&dts Satea 



Isolated 



Paired* 



Grouped 



loa. 



IS 



3 

IIS 



Total 



1X0 



# « Average value of 200 minutes talcen from total observatioQ time of 2U00 
minutes for each ftog. 



•5» ' - 



XABI£ IQJl 



flSEASISnCAL !QSST OF DIFFERENCES AtOTG TEOS I£K>LA!IED^ PAIRED 
AHD GROUPED Sllf lATICMIS COHSDERED IN TAH2 10 



Ccnrparlsoci 


t 


P< 




Isolated -Paired 


0.k2 


0.7 




Isolated-Grouped 


3.9T 


0.01 




Paired-Grouped 


2.69 


0.05 





TABOE U 



nUHBSR OF a?IMES EACH TBOG OBTAIMD FOOD FIRST WHE MBQB OP 
A PAIR - EXPERIMBHTS I MP 11 - AND WHILE IN A GROUP OF THEL7S 

SXFXRIMDiT II 



Frog 



I 
I 
I 

9 

10 

u 

12 



Paired 



Frog 

101 
102 
103 
10i» 
105 
106 
lOT 
108 
109 

HQ 

112 



10 
13 
9 
8 
U 
10 

17 
Ik 

15 
6 

3 
Ik 



6 
i» 
3 
7 

6 

3 
0 
1 
0 
« 
# 

u 



Total 



132 



130 



k5 



TABtB 12 

ATEBAGB HlMBaSR OF SECONDS TAKES BY FRO GS WHSTJ IS0LA3EEID 
(TO OSCAIK EACH OF FOOD > EXFSRIXEIfIS I AHD H 



Frog 


Average Tlste 


Frog 


h. 


. . -ii^: ■ . . 


101 


e 


7» 


102 


3 


768 


103 


k 


68 


l(A 


5 


7k - 


105 


6 


157 


106 


I 


1^ 


107 




220 


108 


9 


137 


109 


10 


130 


no 


n 


83 


HI 


12 


118 


112 



Average Time 




lEABES 13 

RAHEDIGS OF FE«X}S IH YARIOOS CAEEGORISS 
SmLRIMESTS X MB 11 



Frog Hip Puah Ajfproaeh Haknown Hip Push 



Grouped Food Food 

Eat«i Tijae Weltftt 



% 

I 
I 
I 

9 

U 



3. 
t 

I 



X 

I 

i 

I 

9 9 

n 20 



10 

9 

i 



U 
T 



1 
i 

s 

5 
9 
10 

u 

8 

6 
7 



EXnOPJMEHT I 



8 


2 




12 


12 




1 
5 


1 




f 


1 

19 




u 


11 




10 


8 


9 


1^ 


7 


12 


3 


5 


10 


7 


6 


U 



in 



109 
lU 



i i 

I 
I 

10 u 
U 12 

9 9 



t 

I 



6 
2 



7 
5 
1 
l^ 

9 
12 
11 
10 



EXPSRIMEaiT n 

f 

i 

u 

12 
10 

I 



5 
9 

! 

k 

u 

12 
10 
8 
9 



7 
X 

I 
f 
9 

3 

k 

10 

11 
12 

a 



2 

I 

2 

2 

12 

5 
6 

i 



lABlX Ik 



BARE CORBELmORS 07 TSS CMEOOIRZSS VTIB IBSfS IIIDICAIIlia 
PBOBABILm OF CORBELATIOH Mim AS LARCS OR LABGSB 
SUE TO CHANCE AUSM (Edwards, 195k) 



CoQcparlsoaxs 



Frogs 1-12 




from 101 


- 112 


Correlatioa 


P< 


Correlatlm 


P < 




O.QL 


0.8l 


0.01 






.70 


.01 


,ek 






.01 


.ko 






•05 


•32 


.3 


.77 


.€0. 


.IS 


•a 
* 




•Ql 










• • • • 


• • • • 


.71 


.01 


• • • • 




.76 




• • • • 


• • • « 


.97 






.oe 


.85 






.1 


.90 




,6k V 


.09 


.87 


•03. 


.56 


.1 


.76 




.8e 


.03. 


.27 ■ 




.73 


.OL 


.36 




.83 


.01 






.52 : 


.1 


.06 




.52 


^ 


.91 




.67 


.02 


.80 


' m 




,1 


.80 




.08 


.d 


.63 


m 


« .73 


•01 


.26 




.10 


.8 


.70 




.38 


.3 


.28 


.1 



KLp/Puali 
Nip/Appapoada 
Push/Approach 
Nip/ltotaiowa 
Push/lSikncnm 
Approach/UbloioHii 

Paired Grouped y 
ItLp Hip 
Puah Push 

Hp ' 

Grouped Nlp/Ghra>i^d Push 

Nip/FOod Cmsvmtptiaa 
Posh/Food Consiairptiaa 
Approach/Food Cosisisiipticxi 
tfaJmown/Food Coasuntptloa 

Hip/Food Zine 
Push/Food Time ' 
Approach/Food SiiM 
tiaiaxown/Tood !Eba« 

Hip/WBiejxt 

Push/Weig^it 
Approacli/Wei 
Uokaown/Veic^t 



Food Consuniptioai/Wel^t 
Food Slme/Wei^t 



©ISCUSSIOH OF KESUiaS 

'iOils study is coQcexned vlth three basic probLns all dealing vlth 
hierarohial 'bsbxTiors the behavioral display of aggrsssion in fjrogs, indi- 
vidual differences as to aggressive behavior and factors related to indi- 
vidual differences in agsnMisiQti. 

She statistically sigoificant eorrelatiocs that are se«a in flible 
1^ vhen ranMngs of frogs in tlie nip^ push and approach categories are com- 
pared support the assimptiaa that thett |j)^x«e categories represcsxt different 
expressions of the same thing, namely, aggression. In addition, the 
hierarchial standing of frogs 101 - 112 In -Qie nip and push categories as 
indicated tiaa pali^Lngs are higjily cozrelated vlth hierarc hi al positians of 
the MBM frogs \di«a observed in a group of twelve (Tlables 6 and lU). fhus, 
frogs nipping the laost also posh most nbether they are observed in successive 
pairs or as mflsxbers of a group of twelve. 

Correlations between rankings in the uaknoun and aggressive cate- 
gories are generally much lower than correlations among razUdngs in each of 
the aggressive categories (Hkble l^i'). Hhis is evidence tlxat the unlsnown 
category is different and It laay include aggressive activity of a low 

level, it also indtides aotivltieB of a noai-ftggr«ssive nature. 

suable 7 shovs the frequencies of the nip, push, approach and uixknoun 
categories t^en frofs wur* saired. 3Sm unkncwn oatetfary oalEM up about ^0 
per cent of all contaets, ■wifliat ovttr 30 per eent of all eoataets shov a 
veak foim of aggression referred to as the approach, a little more than 10 
per cent shov definite aggression in the push and sonetdiat less than 10 per 



emt lndle«t« the aost IntaaiM farm of aggressloa, namely, the nip. Sables 
6 and 8 give frequencies of the nip and push oategories vhen frogs vere in 
a grcn;^ of twelve* Tbm relative frequoicles of nips end pushes are of the 
MM order ^diether animals are paired or In a group of twelve. A negative 
relationship exists between intensity of aggression and its freq^ieney of 
eapresslon i^ther trogfi are observed in a series of pairings or in a groi^ 
of twelve. . 

Each group of firogs dlcqplaysd a rather loose foim of hierarchy based 
tQKBx differences ooDcag individuals in aggressive pot«itlal as measured by 
the nip, push and approach. Sables 2-3 show the structure, arrangonent 
•ad extsnd of these individual differences. Wide variaUons among frogs 
are Indieated y/hm all aggresslTS aetirity is totaled for each amlnal. 
XUfferenoes in aggressive activity between individuals in any one pair are 
noted by cooqpering totalis for each grog within the chart* flftgr pcx* cent of 
the differttxces between members of pairs are statistically significant as 
indicated by asterisks in the squares of the greater totals. Ttae per cent 
of sl0ilfleaatly dlfferwit totals remains consistent for both groups of 
frogs tested and also for the thzve aggressive categories. 

Bank oorrelation between total aggressive activity esipressed and 
total aggressive activity permitted in this study is negative and statistically 
significant. Frogs with the bluest aggressive potential, as measured by 
sudaer of nips, pushes and approaches administered, eaperleooe the tmm9% 
a^resslve advances from other frogs and vice versa. IMs sugsssts IStat 
aggression Is a distinct foxm of behavior and is based upon certain behavioral 
cues not detected by the observer. 



c-;,.,;' ' Oi^er iafoxnatloti recorded during periods of Q^wrmfeiais In c lud e d 
the follotfiug: (l) wei^ts, (2) amounts of food eaten aaid coBrparisons vh«i 
fJrogs imre isolated, paired and grouped, (3) time taken to obtain food 
virile iscdbted and (k) nximber of times food v&s obtained first vhen indi" 
viduals were in pairs or in a grovq? of twelve. . ^. 

TablA Ik aHaam correlatloas between all of the faxrbors. It Is 
trca -tiwse eorrelatlaas, Ixued upoa rankings of the frogs (Table that 
most interpretations regarding relatlonsJaipe are made. • < 

" t Weigjit dlffereoces sd^t be thou^t of as one ejigpressloa of physio- 
logical variation. When rankings in wel^t, obtained fraa Zeble 1, are 
correlated with rankings in the nip, push, approach and imknown categories 
the resvdts are variable (Skble 1^}. Differences noted previously betwMm 
aggressive bdsavlor and the unknown category carry throu^ whezi the wei^t 
factor is considered. Four of the six correlations between the aggressive 
i«tgtgoirles and wel^t are statistically sl^ilfleant and the othar two are 
almost 80. All are positive. Velght is considered to be a factor la 
bierarchlal behavior of Umam frogs and heavier froigs tend to rank hlgber 
in the three aggressive categories, niBMly, vlp, puih. and approach, yet do 
not necessarily rank higgler in the uzxknown category. 

ffehles 9 and 9A show how frogs rank In individual food caisuDptlaa. 
differences in food consvpption are not as clear-cut as those within each 
of the aggressive categories. Less than ^0 per cent of the pairings show 
significant differences between frogs as Indicated by asterisks within the 
tables. Also, tAien rankings of frog? in total food consvaoed (totals in 
bottoa row) are caqpared to ranldnga in total food coasuned by others in 



their preseace (totals In rl^t coltnm) Mbe results are Inconclusive. Group 
I shows a slgEdfleantly hlgjti eoarrelatloaa, Grotrp II does not, althouf^, all 
correlations are negative. 

, : ^ ISmq renMngs in vel^t are ccccpared to rankings in aiaount of food 
eaten the results are variable. ^Bm heaviest animals In Group II tend to 
•ftt Urn most vherefts the heanrlest iu Qrouip I do not neeasflarlly •at tb* aost. 
^hen ranMngs In tooA emstnptlQn are eorrelstttd tdlb thyote In three 
aggressive categories and the unlmown the results are statlstiC6Llly signif- 
icant at the 5 per cent level In six out of el^t cases and close In the 
raaalnlng two. dere Is no T>reak In degree of correlation observed here, aa 
In previous cocrparlsans, "between the three aggressive categories and the 
vcbkamti, ■which indicates the more aggressive as well as the niore active 
frogs eat more than the less aggressive and less active animals. 

Sie srezase tljae ta]fiea l>y isolaiied aaiOAls to Iocftt« food varies 
vldely anong inttvlduala (Wble 12). Qmm dlffWrwices May ^ to 
variations In the chemoreceptor systems and in ability to respond to -ttiese 
stimuli. When rankings in this category are cooipared to rankings in the 
aggressive categories (Table ill-) the correlatians, while all positive and 
significant at the 5 per cent level In Orovq} I, are positive "but of 
questloaaa'ble sigalflcenee in Group H. When rankings in aioount of time 
taken to locate food are ccccpared to rankings In amount of food consmed 
th« coi7elation« are not consistently hi£^. Sh^refore, the xoore aggressive 
firogs do not neeesswrlly find fbod more galekly, nor do the frogs eating 
more while paired necessarily take the least time to locate food \iben. iso- 
lated, t.; ■ , . . ■■■ 



IxifQaxmtim aa the nusber of tljaes each fjrog obtained food first 
>)btle a noftier of a pair and vhlle In a grotq) of tvel'V« Is gtrsa In Table 
U. Whidx animal obtains food first at the start of each series of ohser- 
vatloos is apparently a chance phenorattion not noticeably related to any 
factors previously discussed. . 'v • - 

It Is possllale that the three aggressive categories represent 
attempts by frogs to eat one another. Cannibalism Is veil knona In this 
species especially -whan size differences are great and availability of food 
Is limited (Rose, 1950). 2he behavior resulting in a alp vas observed to 
be very sladlar or Identical to that leading to constnspticEi of Uver. Osily 
the size and movement of the other frog semed to prevent the coanpletion 
of the act of eating In the case of the nip. Ihe Increase in frequencies 
of nips, pushes and approaches in the presaice of food is further evidence 
sufg^orting this asstsiptloii* If the observations noted above are valid the 
atteorpt to obtain food laay censtltute a camnoa basis for aggression in laaagr 
vertebrates. ;•. ,• ■ ■ . 

;'; A short tis» afber a tult of food Is plao«4 in an a<fiBriua vith a 
group of twelve frogs the frequency of sweeping motlctts with front Iteert and 
the amount of sulusnlng activity both increase as do frequencies of all types 
of contacts. Hvrnx after food has been eaten the Increased activity caatinues 
for 10-30 minutes. A searching type of behavior by one or two frogs in a 
grovg? initiates the same sort of activity afflong the otlicrs even in the 
absence of food. Sms, aetlvlty in the group can be Initiated by the behavloar 
of one or two members, and this factor enhances the chances of locating food. 
79^d ftLfiO la found a twi ^ftr by ^harnw vlth siora tiIdmiI b ooGuj>|yis^ a oMrtaln 



roluae of vater, mA In adflltion iSbm greater amount of Infiiviaaal activity 
no doUbt requires tbe use of more energy. Tiahle 10 shows the compjurative 
aoounts of food eaten by frogs viaea isolated, vbesa. paired and •whan In a 
group of twelve. Hhe total amount of food consumed by frogs varies directly 
Kith the nuniber of animals plafted togetlxir. Differences in total food con- 
suinpUm betweei the isolated and pairiA iltuatians are not staUstically 
Biffiificant, but tihen ftogs are observed in groups of twelve the total con- 
sanption is slgilflcanUy greater than in either of the other two situaUons. 
If increased food cansumptlcn is actually of poslttve value to the anJaal, 
this may be an esaiQtle of cooperation or the Allee Effect (Odum, 1953) • 

•Che present work has indicated the existence of certain b^wviaral 
«itlties by the method of analysis. It has also attcaapted to relate these 
entities one to aaother and relate thsm to certain broader aspects of 
living things ouefe as iihyelology, agg»»iim, copperaUon and hlerarchlal 
behavior. An atteimrt at controlled observation is used here to facilitate 
an mderstandlng of behavior both In one species and as a ganeral phenomenon 
of life. While one person has been studying the brtiavlor of these frogti 
the data as tabulated In a asnse have reflected certain aspects of human 
behavior. On the whole, correlations are hlijier in (kna^ II taaan they are 
in Gxoiq* I, when rankings in the three aggressive categories and -ttie unknow 
«re coogpared to each other and to rankings in other factors. It could be 
that frogs in Qmapa 1 and II vere behavlorally dlffenmtj on the other hand, 
as a result of continued work vlth the animals, the experimenter may have 
learned to fit certain aspecte of behavior into categcxries with greater ease 
in the second groi^). Assuaalng iatie latter, this probably represents greater 



accuracy tn dstexminlng wt^efseSjta In Qroup II. Hofverar^ there still is a 
source of danger in the metbod of analysis used in this study. Blvlslcsi of 
the bdiKvioral yboHA Into gui^ increasingly aGn>Bx«at categories or entities 
■i^t eneeniMgi m eibeerver to igaore relatiaBdilp? between peu^s and the 
bduYioral vfaole. Cie parts versus the vhole dichotomy is one that science 
luMi AM9ed £rm the ,pt«rt, because an inqportant feature of the scientific 
ttetliod is the analysis of vholes into paarts. Often diehotaoies are eliffl* 
Inated as more is known about a subject. For Instance, the study of heredi-ty 
moA suTiroament has becosie not so imich a pcroblen of two separate things as 
different aspects of the same thing. ItLkeviBe, future vork may shov that 
hairarehial behavior is a special case of territoriality or vice versa. 

, . Mooh of this vort: bai e sn temd around liw study of tadlvldual dif « 
ferences in aggression, i. e. the determination of the presence of and form 
Of biararchial }»tBmriM iA this £rog. An analysis of hierarchlal behavior is 
also attenipted. An increasing number of studies In animal behavior are not 
concerned vlth an analysis of both the phenomenon of aggression and of the 
hierairohy (Allee, Foreoan, ftuiks and Holablrd, 19?^; Braddodc, 19^(91 
Bovjberg, 1956 j Crawford, 19l»0i Colllas, 19^3i Douglis, 19^) Gre«iberg, 19^7) 
and Delany, 1955)* Such studies njjmamt att«spts to analyze the broader 

aspects Of behavior. . - 

Many European workers, especially theebhologlsts, emphasize the 
miilywLa of more basic betuvrioral eatitios m is evident in the studies of 
releaser mechanisms (Tinbergen, 19^^^ 19^^ 19?3), social faeilitation 
(Barling, 1938j Davles, 1953)^ displacement activities (Lorenz, 19^1j 
Wnbergen, 1953) and imprinting (lorenz, 1952K . ; , 



Sbie cLLff erences vhlch exist betveea these tvo schools iu their 
mpfproaches to the geaeral study of behavior are also evident in their 
^fffffcmahies to the study of bdiavloral evolutioa. Sils is clearly shovn 
tilum the Su3^^pean viewpoint discussed by tOEtRiz (19^0) Is coopered with the 
American airproach as Indicated by Allee (1952). ^Ebe Europeans attempt to 
coorpare oore basic bn^Msviorttl entities aiacxig dif£to«&t «gp«ol«8 KiiyBreas the 
Americans attenq^t a ccepearison of more general phenoBMna. Both astproachas 
are of course useful in any study of evolutiacL be it organic or behayloral. 



A study «M mdi of two groups of twelve frogs In an ftttesapt to 
amalyie «sgrossicin and to drtemLne ^Aetlier inOividaaU wlIMn the gjroi^ 
y [lY ^an iggressive differeaces resultlxig in hlerarchial behavioor. 

1» Aggressioo in Xgoopus laevis Tln^^^n aiJ^Ojaditid ■behavioraUy ia one 
of three ways, namely, "by the ajjproach, tflie pttflh sod the nip. These 
Obtteories also represeat iaoreasing intensities of aggressive behavior 
in tSie oraer given. 

2* Contacts repires«iting the aore intense categories of aggresslTe 
1»^hearlor occur le«9 ftsfMatiy ttm do tHoee r«(pres«atin6 Hio less inteoae 
ones in 'bath the paired and grcuped situatione. 

3. Large differences aoang frogs are indicated vhen aggressive 
ecfcivity is totalled for each anlaal. Progs displaying 1^ aost aggressive 
activity in one category tend to display the aost In the others* 

k» Differences in «ount of aggressive "behavior displayed betwMtt 
airftan of pairs are statistically slgplfloant in ^ per cent of the cases. 
!Ihls is ccQsistent for each of Ha* aggressive categories. 

ft A category '^ris pn*^ as unkoown repres«a.ts contacts in "which, no 
a^resslve S Ufxm &m MmMI mOmXa could be detendned. She total 
frequency of these kinds of contacts is as great as all other types cooibined. 

6. Frogs vlth greater aggressive yoteatials, M Msured by nuaber of 
nips, pushes and approaches, experieaace fewer aggressive advances flrom otiier 
fro^ and vice versa. 

7« Seavler frogs display aggressive behavior aoore frequoatly than 



lighter caaes, yet are xxot necessarily 'tiM aore active anlasls (aotivity 
referrin^ to raoMixg in the untooua catee^ry). 

8. Sbe more aggressive as imU as oore active £rog& eat more thaa the 
lees aggressive and less active anlneifl, ytt lAiiii renMngs in total food 
coasuoed are compered to raaldngs In total food consumed by others In their 
presence tiie results are sot necessarily Tal^oly correlated* 

9. K> cousistoit relatlonehlps exist amoig rankinfls in fdod can- 
suB^tioti, tim l&ken. to locate food viiile Isolated and vel^t* 

IQ, As the nualber of anljoaals placed togttixer is increased to twelve, 
the amount of food ccxisuaed "by each troQ increases. !Ihls may "be an eTB Mi ila 
of cooperation or the AUm fiffeot* 

XI* It is possl^ thttt aggresiiiit« 'behavior in these fSrogs reflevto 
an attatrpt to eat (me anotberj as "behavior resulting in a nip is nearly 
identical to that vhich leads to ooiisuBQitim of liver* 



Allee, V. C, 1951. Cooperatlm amaag AnlTnols, vi-fch Hvinan SgpHcatloos. 
IfeifwwOT, Tcnrk. 233 PP* 

— , 1952. Dnmi nance and Merar<day In societies of verteTsrates; cited 
in Stsnicture et Hiyslologj' des Soc?.etes Aolnales, ed. Pierre Qrasse, 
( Coll. Ihtemat. Centre* Bat. Beah* Sd.i ^} 157-183- ). 

and J* C. UcMnson^ 195^> Dominance and eiibordlnatlQQ. In the 
amfoHi. aogflj3h, Mustelus cauls (mtchill). Pfar^'Biol. Zool., ^{k)t3^'-3l6h. 



— — , and M. Doviglla, 19^5* A dotalnaace order in the hermit crab, 
fliygaa IfUfi^ipHDW aey. Bcology, ^tkllAlZ* 

Focrenen, E. M. Banks and C. H. Holabird, 1935* Sffeets of an 

androgen an dominance and siibordlnanco In six ccoman "breeds of Galltts 
ffOlus . Phyalol. Zool., 28(2)1 89-U5. 

B. Gbreenlwrg, a. N. BoMithal and P. f!rank, Sons effeets of 

social orgBnlzatioci on growth In the green simfiah, I«ponls cyanellTia . 
Jour» Eacp» Zool., 108:1-19. 

Banks, E. H., 19^6. Social orggoilzatlca ift Md twl hma [ (hUxm 

gRilus sub^. ), Scolosy. ^i239-2.kQ, 

Bennett, M. A., 19^* Tbe social hlerardiy In ring donres. Ecology, 20(3)< 
337-357. 

Bovjberg, R. V. , 1953. ilcnlnanoe co-dar In Hm en^fiah, Qronectes vlrllla 
Hagsn. fhyslol. Zool., SjSiXl^l^ 

1956. ScDMi factors affecting a^resslre "behaivlar In cra^ah. 
Phyalol. Zool., ^:127-136. 

Biraddock, J. C, 19i^5. Soma aspects of the dnmlnanee-st&ordtnatlon 
relationship in the fish, Platypoecllus aaculatns . Riyslol. Zool., 
2^:176-195. 

, 19'^9> Bie effect of prior residence vipcai danlnanee In HtM flah, 
Platypoecilus aaculatus . Physiol. Zool., 22:l6l-l69. 

Carpenter, C. B., 19l|-2. Sexual behavior of free ranging Rhesus iBODkeys 

( Maeaca aulatta ). I. Speelmms, procedures and Isehavloral eharaeteriatloa 
of estrus. J. Comp. Psychol., 33;113-1^>3* 

ko " i 



eolilasj Ifi 1., 19^3. Statltstleal aaalywls of f&ctors -Hfalch make for sixccew 
in Intitlal «icount«r8 Txitween hens. Aa« Hat»« 21: 519-5 38« 

— 19Mt^. AggresBlve behavior amoog vertebrate animals. Phyalol* Zool«> 
1X:83-123. 

Crawford, M. P., 19^^0. Tbe relation between social doalnance and the 

BMnstrual eyele tn fesoule chl^panzeea. J. Ckanp* Peyebol*, 30:1*83-513. 

1911^2* DcBdnanee and social behavior, for chimpanzees, in a nm 
competative situation. J. Cflpp. Psychol., 321:267-277. 

Darling, F, P., I938. Bird Flocks and the Breeains Cycle . 2he Iftilverslty 
Press, Cafflbridge. 12^4- pp. 

Daudln, F. M., I803. Historie Ifaturelle des Rainettes. des Grenouilles et 
d6S Crapauda . Paris, p. Q5» 

Davies, J. L., 1953. Colony size and rao?roduction in the grey seal. Piroo. 
Zool. 80c. Londfln., 123(ll): 327-332. 

Delaney, M. J., 1955. ^ erolution of aggressive behavior in vertebrate 

animals (tMpiibllahed). Dept. of Zool., Glasgov, Scotland. (How at 
Ifetiv.of Southharaptcn, England). 

Dlebschlag, E., 19^1. Psychological observations on the rank order in the 
domestic pigeon. From the Zool. Inst, of the Ifelv. of Marburg. 
(PsychologLsche beobachtungsn uber die xtuigrardnimg bei der haiistaube. 
Zeitachr. f. Tlerpsychol.^ 4:(H.2) 173-188). 

DougLis, M., 191*6. Seme evidence of a flomlnnnce -subordinanee relationship 
onang lobsters. Anat. Rec., 2i'553» (Abstract). 

— — , 191*8. Social factors inflvienclng the hierarchies of aoall flx>ck8 of 
dOB«stic hen: lha interaction between residtat and part time loembers of 
or^tfiized flocks. Physiol. Zool., a:ll*7-l8a« 

Edwards, A. L., 1951*. Statistical Methods for the Behavioral Sciences. 
Rhinehart and Coorpany, Inc., Binr York. 5^*2 pp. 

Bllan, E., and R. W. Murray, 1951. New laterlal line sensory organs in 
Xenopus laevis. Nature, 168(1*272:1*77* 

Ck>in, Coleman J., 1953. Parsonal Conanunication. -^''^ 

Oola, Olive B., 1955 . World Outside >ty Poor. MacMlllaa Co., New York. p. 1*3. 

Gordon, K., 19l*0. Territorial behavior and social dooinance asiong Sculridae. 
J. Mwwaal.. 17:171-172» . . 



Grant, W. 1955 • •Mtrrltorlallam in two species of salaajaad^s. Science, 
1S:(33.35)J137» 

Oreenberg, B., 191*7. Some relations between territory, social hierarchy 

and leadership in the greeri suailsli, Lepotnls eyan^OJLus * Riysiol. Zool« 
^: 267-299. 

— — , and G. K. Koible, IS^th, Social behavior of the Aa»rloan chameleon, 

Anolis carol jnensls Volgb. Physiol. Zool». 17:-^2-4^, 

Oi&l, A. M., and ¥. C. Allee, 19hh, Seme measureable effects of social 
organization In nocks of hens, fhyglol. Zool.. 18:320-31*7. 

BsnMcrstroni, F., 191^2. Sondnanee in vlnter flocks of chickadees. Wilscn 
Bull. , 54:32-1*2. 

Hasklns, CP., 1951 • Of Societies and Mai . Horton, ITev York. 2^ pp. 

Edward, W. E., and J. T. Salem, Jr., 19l»2. Intercovey social relationships 
in the valley quail. VUson BuU., §4:162-170. 

Kslth, A., 191*9. A .Tfev !Iheory of Hianan Eyolutlon. Philosophical Ubrary, 
New York, 1*51 pp. • ■ ; : ;.. - . - 

lAslle, J. M., 1890. Notes on the habits and oviposition of Xaiopus laevis« 
Froc. Zool. Soc.. London. 69-7I. 

lorenz, K., 191*1. V«*glelohttide bewegungsstudlen an anatinon'. Jour. f«' 

Oroithol.. 82(Pestschrlft Belnroth):19l*-29l*, 

——-1 1950. The conparative method in studying innate behavior patterns. 
Physiological Mechanisms in Animal Behavior . Acadsmic Press, New York. 
221-268. 

— — , 1952. King SolcEm's Ring . Ihoinas Y. Growell (Jo., New York. 202 pp. 

lorn, M. S., 1956. Dominance-subordination relationships in the crawfish, 
CambareU.us shufeldtli. Tulane Studies in Zoology. 1*(5):1^-170. 

Martof, B. S., 1953' Tisrrltorlallty in the greeox fSrog, Rana clnwltj^nw . 

Ecology^ 2it'l56-174. 

Maslow, A. H., 19l*0. Doodnance qfuality and social behavior in infra -human 
primates. J. Soc. Psychol., 11:313-321*. 

Munro, A. F., 1953* ^e anmonia and urea exeretion of dlffer«it species of 
amphibians during develosment and metaBorphosls. Biochem. Jour.. 
5l*:29-36. . i '- ' 



Wmmei, M. a., 1956. social "behavior and Inter^ciflc canrpttltlon In two 
trout species. HiyaiQl. Zool.j 22,^611-81. 

Odbaa, I. P., 1953. FuoaacientalB of Icology* V* »• Semders Co., Philad«lpiil*. 

Pardl, t., 191*8. DoBiiaance order in PolisteB VAsps. Physiol. Zool., gLtl-lS* 

Pirker, F. Jr., S. L. Kobins sad A, LaTreriage, 19**?. Breedijas, rearlag and 
care of the South African clawed frog (Xeoogjs laevls) . Baa Aawr. Mat*, 

ja: 38-^9. 

PBiteraoa, Bellie P., 1951. aasal cavities of "ttbe toed Haaiplpe caryalhoi 
Mlr-ril) and otiber Plpidae. Proc. Zool. 80c., Loadoaa, l^t2JJ3ai-iH5. 

Peargon, P. G., 1955 . Population ecology of the Bpedefoot toad, Seaphiopus 
\ Hblbroold. (Harlan). Ecological Monoscanhs, 2^:233-207. 

Bose, W., 3^0. The Reptiles and Actplii'biana of Southera Africa. HMfcMr 
Miller Ltd., Cape Town. pp. 1-3^. 

Sabine, W. S., 191*9. Doainance in vinter floclcs of Juncos and tree ^perrowa. 

Riysiol. Zool., ^:61v-85. 

Sdijeiaerup-EbTae, T., 1913. Honsenes stsDBie fidrag til Honsanes peyiwlogl 
aatiaea, 2L»2^-276. 

— — * 19^» Social behavior of birdsj cited In C. A. Murchlnaon, A aandbook 
of Social Psyciioiogy. Worchester. Pp. 9^7-972. 

Sharif, M., 1956. Experimeats in ffccnxp cooflict. Scientific American, 
121(5) »5^-58. 

l^st, F. H«, 195l». Social aegresslveneas in an aacEMbian. Sci«ice, 
120(3100) »1^> 

Tittbergan, H., 19lt2. An ob.iectivi3tic study of the innate beba-/iour of aniaals. 
Series D. Bibllothica Biothcoretica. V.l pars 2. 1. J. Brill, london. 
pp. 39-98. 

19li8. Social releasers and the experimental xaethod required for their 
study. Wilson Bull., 60:6-52. 

1953. Social Behaviour in Animals, vith Special Beferences to 
Vertebrates . Methuen and Co., Ltd., Lcndoi. J<^ Wiley and Soas, Inc., 
liev York. 150 pp. 

Uhrl6h, J., 1938. The social hlerawShy In aXMno mice. Jbur. Cocg). Faychol., 
g5.»273-^3. 



MetrogravTge. BBltiaiore^ Ud^p ApriLL ^i* 

Ytix^e, R. NU, 1939* Social <3anlnanoe and seaenaT itatns in the nMnrpan: 
ghe Quarterly Beviev of Biology. l^t»115<436« 



BIOGSRAPHICAIi X9SM8 

Bobert Bice Baubrlcli vas Ixxax In CilscraBiaat, Sew VSaxagOilxtif Hay k, 
19S% aad gracbuaiwd txm St«r«u EL^ fi c h o o l ^ GLaxmaeat, ija. June, 19^. H» 
catered the t]Dlv«r8l% ot Wn Vtupi&sILn in iSm ftiSX of 19^ and after thrott 
MMsters entered the Axny Air Force lAiere he sp«it three years. 

In ibrch^ 191^, he entered ItLchlgaa 8tate IMTer^i^ and receiwA 
a Bachelor of Science degree in Forestry (l9*^9) and a Master of Science 
degree In Zcxtlogy (19^2) £roa that inatituticn. Ihe sisimer of 19^2, iias 
spent at Point Barrov, ilLaska, vith the Office of Iforal BesMxtfli. He \m§m 
his dcxrtoral program at the IMversity of KLorlda in September, I952 and spent 
the ■ini II of 19^3 and 19^ at the ifarine Biological Laboratory, Mooda Hole, 
ilHHnchusetts. VhUe at the Oiiversity of florlda he tan£^ biology 
Inlxn-atory and ganeral biology (C-61, C-^} and also vas Assistant Baaiteit 
Adrlsor In one of the housing areas on canqpus* 

Haubrloh Is a neaber of XL Si0Mi fl, Rd BL^m, Blgna XI, fhe 
Association, foor the Study of Aoifflal Behaviour and the Ecological Society 
of Aaerloa* 



ahls dlasortatioQ "was prepared under the direction of the c hairm a n 
of -aift candidate ' s supervisory committee and has "be«i approved by all 
msmberB of the conunittee. It was suhmitted to the Dean of the College of 
Arts and Sciences and to the Graduate Cornell and vaa apjaroved as partial 
fulfillaent of the reqvUreoi^its for the degree of Doctor of Philosophy. 



Au^ist 10, 1957 



i-'<' 



"Smmf CkxUefte of Arts azid Sciences 



Been, Graduate School 



SUHSR7IS0KY